Jean-Frédéric Terral

Primary domestication and early uses of the emblematic olive tree: palaeobotanical, historical and molecular evidence from the Middle East

Our knowledge of the origins of olive tree domestication in the Middle East and on the processes governing its extension and persistence in different vegetation types from prehistory through antiquity to modern times derives from diverse sources, spanning the biological sciences to the humanities. Nonetheless, it lacks a robust overview that may lead to floating interpretations. This is especially true in the Middle East, considered as the cradle of agriculture, and where the evolutionary history of this emblematic tree is intertwined with that of civilizations. Olive fruit, oil and wood have been, since Prehistoric times, characteristic products of the lands bordering the Mediterranean Sea. In the domestic economy of these countries, the olive tree gradually became a traditional tree crop since the first oil extraction, through the emergence of regional commerce that accompanied the rise and fall of early Near‐Middle Eastern urbanism, until the development of modern trade, with an oil production estimated at circa 3000000 tons per year. The rising importance of the olive tree in human life has turned the tree into an endless source of fascination in the Aegean and Eastern Mediterranean, a symbol and a sacred tree, widely cited in the Bibles, the Koran, and in ancient literature. Here we argue that advances in radiocarbon chronology, palaeobotany, genetics, and archaeology‐history have profoundly refined the history of olive trees in the Middle East. This review shows that the heartland of primary olive domestication must be enlarged to the Levant and not only focus on the Jordan Valley. The domestication of the olive tree is a long and ongoing process, linked to the early production of oil and the development of the olive trade. We also suggest that the olive tree became a particular icon, a sacred tree, during the Biblical period in the Levant.

The origins of the domestication of the olive tree

The present diversity of the olive (crop) and oleaster (wild) tree was investigated with nuclear and cytoplasm markers. Patterns of diversity of the wild form inferred eleven ancestral populations in the East and the West of the Mediterranean basin. Patterns of diversity for cultivars are less clear, but we showed that cultivars admixed to nine groups that corresponded to oleaster ancestral populations. We inferred that nine domestication events took place in the olive, but these origins were blurred by gene flow from oleaster and by human displacements. These origins of domestication probably reflected different reasons and uses to domesticate the oleaster.

A small XY chromosomal region explains sex determination in wild dioecious V. vinifera and the reversal to hermaphroditism in domesticated grapevines

BackgroundIn Vitis vinifera L., domestication induced a dramatic change in flower morphology: the wild sylvestris subspecies is dioecious while hermaphroditism is largely predominant in the domesticated subsp. V. v. vinifera. The characterisation of polymorphisms in genes underlying the sex-determining chromosomal region may help clarify the history of domestication in grapevine and the evolution of sex chromosomes in plants. In the genus Vitis, sex determination is putatively controlled by one major locus with three alleles, male M, hermaphrodite H and female F, with an allelic dominance M > H > F. Previous genetic studies located the sex locus on chromosome 2. We used DNA polymorphisms of geographically diverse V. vinifera genotypes to confirm the position of this locus, to characterise the genetic diversity and traces of selection in candidate genes, and to explore the origin of hermaphroditism.ResultsIn V. v. sylvestris, a sex-determining region of 154.8 kb, also present in other Vitis species, spans less than 1% of chromosome 2. It displays haplotype diversity, linkage disequilibrium and differentiation that typically correspond to a small XY sex-determining region with XY males and XX females. In male alleles, traces of purifying selection were found for a trehalose phosphatase, an exostosin and a WRKY transcription factor, with strikingly low polymorphism levels between distant geographic regions. Both diversity and network analysis revealed that H alleles are more closely related to M than to F alleles.ConclusionsHermaphrodite alleles appear to derive from male alleles of wild grapevines, with successive recombination events allowing import of diversity from the X into the Y chromosomal region and slowing down the expansion of the region into a full heteromorphic chromosome. Our data are consistent with multiple domestication events and show traces of introgression from other Asian Vitis species into the cultivated grapevine gene pool.

Bioarchaeological insights into the process of domestication of grapevine (Vitis vinifera L.) during Roman times in Southern France.

Grapevine (Vitis vinifera), one of the most important fruit species in the Classical Mediterranean world, is thought to have been domesticated first in South-Western Asia, during the Neolithic. However, the domestication process remains largely unknown. Crucial unanswered questions concern the duration of the process (rapid or slow?) and the related geographical area (single or multiple-origins?). Seeds from domesticated grapevine and from its wild ancestor are reported to differ according to shape. Our work aims, first, to confirm this difference and secondly to identify the extent of domestication in the grapes cultivated by Romans in Southern France during the period 50 BCE–500 CE. We had the opportunity to analyze uncharred waterlogged grape pips from 17 archaeological sites. Based on an extended reference sample of modern wild grapevines and cultivars our work shows that both subspecies can be discriminated using simple measurements. The elongation gradient of the pip’s body and stalk may be regarded as an indicator of the strength of the selection pressures undergone by domesticated grapes. Grapevines cultivated during the Roman period included a mix of morphotypes comprising wild, intermediate and moderately selected domesticated forms. Our data point to a relative shift towards more selected types during the Roman period. Domestication of the grapevine appears to have been a slow process. This could result from the recurrent incorporation into cultivation of plants originating from sexual reproduction, when grape cultivation essentially relies on vegetative propagation.

Genetic structure of the date palm (Phoenix dactylifera) in the Old World reveals a strong differentiation between eastern and western populations

BACKGROUND AND AIMS Date palms (Phoenix dactylifera, Arecaceae) are of great economic and ecological value to the oasis agriculture of arid and semi-arid areas. However, despite the availability of a large date palm germplasm spreading from the Atlantic shores to Southern Asia, improvement of the species is being hampered by a lack of information on global genetic diversity and population structure. In order to contribute to the varietal improvement of date palms and to provide new insights on the influence of geographic origins and human activity on the genetic structure of the date palm, this study analysed the diversity of the species. METHODS Genetic diversity levels and population genetic structure were investigated through the genotyping of a collection of 295 date palm accessions ranging from Mauritania to Pakistan using a set of 18 simple sequence repeat (SSR) markers and a plastid minisatellite. KEY RESULTS Using a Bayesian clustering approach, the date palm genotypes can be structured into two different gene pools: the first, termed the Eastern pool, consists of accessions from Asia and Djibouti, whilst the second, termed the Western pool, consists of accessions from Africa. These results confirm the existence of two ancient gene pools that have contributed to the current date palm diversity. The presence of admixed genotypes is also noted, which points at gene flows between eastern and western origins, mostly from east to west, following a human-mediated diffusion of the species. CONCLUSIONS This study assesses the distribution and level of genetic diversity of accessible date palm resources, provides new insights on the geographic origins and genetic history of the cultivated component of this species, and confirms the existence of at least two domestication origins. Furthermore, the strong genetic structure clearly established here is a prerequisite for any breeding programme exploiting the effective polymorphism related to each gene pool.

On the origins and spread of Olea europaea L. (olive) domestication: evidence for shape variation of olive stones at Ugarit, Late Bronze Age, Syria—a window on the Mediterranean Basin and on the westward diffusion of olive varieties

Charred archaeological stones of Olea europaea L. (olive) from Late Bronze Age Ugarit, Syria, were analyzed with geometric morphometry and compared with a morphological differentiation model established on the basis of analyses of modern spontaneous (uncultivated) olive populations and cultivated varieties of various origins within the Mediterranean Basin. The results allow a reinterpretation of the east–west morphological diversity previously observed in wild olives. The archaeobotanical data were compared in detail to the partly geographically structured modern morphological diversity of the cultivated olive. Ancient morphotypes could be distinguished, among which one is dominant in the assemblage. Their diffusion from east to west is shown, and their time of arrival in the northwestern Mediterranean can be evaluated by comparison to archaeological material from that area. Combining morphometric and genetic data, modern reference and archaeological material also guides us in understanding the mechanisms that prevailed in the long-term agrobiodiversity of the olive.

The egyptian olive (Olea europaea subsp. europaea) in the later first millennium BC : origins and history using the morphometric analysis of olive stones

The authors examine a sample of olive stones from Egyptian contexts and show that from the first millennium BC, if not before, some of them relate to cultivars originating from the Levant. But equally prominent and just as early is another variety, of unknown origin and currently peculiar to Egypt. The method used is geometrical morphometric analysis – essentially classifying the olive stones by their shape.

Wild and cultivated olive (Olea europaea L.): a new approach to an old problem using inorganic analyses of modern wood and archaeological charcoal

Abstract The elemental composition of wood varies according to the genotype and ecological conditions of plant development. Inorganic elements of wood are preserved in charcoal. Elemental composition analyses of olive wood combined with a multivariate statistical treatment show that wild and cultivated trees and their bioclimatic environments can be distinguished (thermomediterranean and mesomediterranean stages). The elemental composition of some olive charcoal from two archaeological sites in Spain is analysed. The bioclimatic and taxonomic information obtained from archaeological charcoal analyses records the anthropogenic processes interpreted as the transition from a mesophilous phase (Cardial Neolithic) to a thermophilous phase (Roman Age), correlated with the gradual replacement of the former mixed oak forest by the “matorral”. Finally, these results address the question of whether the cultivation of olive trees had occurred in Spain by the Neolithic.

Inferring the agrobiodiversity of Vitis vinifera L. (grapevine) in ancient Greece by comparative shape analysis of archaeological and modern seeds

The origins and biogeographical history of Vitis vinifera L. (domesticated grapevine) remain largely unknown. Shape and size have long been used as criteria to distinguish between wild and domesticated grape pips. Here we have analyzed variations of seed morphology in order to provide accurate criteria for the discrimination of different groups of varieties. Diversity in present-day cultivars and wild grapevines of Greek and east Mediterranean origin in relation to other Asiatic and European varieties and wild grapevines provides the basis for our analysis, which aims to allow the characterization of the ancient diversity of cultivated grapes in relation to present-day cultivars. Geometric morphometric analyses (Elliptic Fourier Transform method) have been used to characterize the seed shape and size of modern and archaeological material using 40 variables per seed. 197 archaeological grape pips from the 7th century bc sanctuary of Hera in Samos, Greece were compared with an extended reference collection of 269 modern cultivars and 83 wild populations, 10,518 seeds in total. Our study confirms the relationships between seed shape and domestication. Modern diversity is partly structured by the geographical origin of cultivars, but influence of other factors may play a significant role in clustering. The wide diversity of varieties offered at the Heraion of Samos during the Archaic Period, including cultivars growing on the island, imported grapes and wild morphotypes, is related to the history and geographical location of the island as well as to the diversity in the geographical range of pilgrims making offerings to the sanctuary.

Holocene vegetation changes in the northwestern Mediterranean: new palaeoecological data from charcoal analysis and quantitative eco-anatomy

The stratigraphic sequence of Montou (Eastern Pyrenees, France) covers a period from the middle Neolithic to the late Bronze Age. Environmental changes are noticed since the middle Neolithic, when the decline of the deciduous oak favours, in the long term, the evergreen oaks. The Chalcolithic period witnesses the arrival and spread of mesomediterranean and thermomediterranean plant formations. This evolution may result from increasing anthropogenic pressure as shown by the maintenance of a garrigue vegetation during the Bronze Age. The association charcoal analysis/quantitative anatomy offers new perspectives concerning vegetation changes. Both approaches record the existence of mesomediterranean bioclimatic conditions during the middle Neolithic; thermomediterranean affinities are recorded first, by charcoal analysis, during the early Bronze Age, and by eco-anatomy during the transition from the middle to the late Bronze Age. Quantitative anatomy also pinpoints (1) the existence of two humid phases, (2) an increase in mean annual temperature, and (3) the exploitation and management of the olive tree since the early Bronze Age.